Management of Amphibians, Abstract.-The primary xeric pinelands of peninsu- lar Florida are lon~leaf pinelturkey oak sandhills and

Reptiles, and Small Mammals sand pine scrub, Their management on public lands is largely confined to prescribed burning to maintain

in Xeric Pinelands of fire climax status of the vegetation. The regulation of

Peninsular Florida1 large-scale developments on private land has stim u- lated interest in preserve design and management. The suite of techniques used to solve conflict be- tween natural system preservation and develop-

I. Jack S t ~ u t , ~ Donald R. Ri~hardson,~ and ment includes: (I) conservation set asides (pre- Richard E. Roberts4

Xeric pinelands seem incongruent with reference to Florida, a state with annual rainfall that ranges from 50-65 in (1 9.6-25.6 cm). None theless, the Florida peninsula contains thousands of acres of sandy soil derived from marine deposits dating to the Pleisto- cene (White 1970). Two distinct plant associations, longleaf pine (Pinus palustris)/ turkey oak (Quercus laevis) sandhill and sand pine scrub (Pinus clausa), have developed on these nu- trient deficient and excessively well- drained soils. Significant areas of these plant associations occur at higher, albeit modest, elevations rela- tive to the surrounding landscape. In fact, certain topographic features, e.g., the Lake Wales Ridge and the Marion Upland, were likely to have been true islands during interglacial periods while the remainder of Flor- ida was covered by a shallow sea. Regardless of their exact origin, xeric pinelands support many relatively unusual species of amphibians, rep- tiles, and small mammals.

'Paper presented at symposium, Man- agement of Amphibians, Reptiles, and Small Mammals in North America. (Flag- staff, AZ. July 79-2 1, 1988).

21. Jack Stout is Professor of Biological Sciences, Department of Biological Sci- ences, University of Central Florida, Box 250CO, Orlando, FL 328 16-0368.

3Donald R. Richardson is Adjunct Profes- sor, Department of Biology, University of South Florida, Tampa, FL 33620.

4Richard E. Roberfs is Biologist, Division of Recreation and Parks, Florida Department of Natural Resources, Hobe Sound, FL 33455.

serves) on site; (2) habitat restoration; (3) purchase and dedication of off-site preserves; (4) species relo- cation; and (5) wildlife resource mitigation fund.

Human population growth (3.3% per year) and development in Florida continues to encroach on upland habitats and particularly on xeric pi- nelands. Most of the habitat loss is to agricultural uses, principally citrus. Oddly, the state's excellent wetlands protection acts have forced develop- ment into the uplands. Thus, xeric pinelands and their narrowly adapted fauna and flora are increas- ingly threatened by area reduction, fragmentation and isolation.

It is our intent to discuss the man- agement of these xeric pinelands in general and, more specifically, in the context of small preserves in an oth- erwise developed landscape. Man- agement of xeric pinelands as ecosys- tems is yet in its infancy, and more detailed prescriptions for designated species are unproven. However, progress is being made (Cox et al. 1987) and improvement and revision of current thinking on management practices is anticipated. This paper summarizes selected literature on xeric pineland and the species associ- ated with these communities to as- sess management practices. In addi- tion, unpublished information has been used and identified in the text.

Preserve design efforts by us have been on behalf of developers re- sponding to development orders pre- pared by governmental agencies. These designs are site specific in de- tail, but nonetheless point to general problems and solutions. Our ap- proach has been to focus on provid-

ing the area required to support minimum viable populations of "keystone" or otherwise critical ani- mal species of a given xeric pineland. Once this area is settled on, manage- ment should focus on those species whose minimum area requirements are met, whereas no special efforts are expended on species with larger area requirements.

XERIC PINELAND HABITATS

Longleaf Pinepurkey Oak Sandhills

The longleaf pine/ turkey oak sandhill association (LLP/TO) was about 15% (2,110,256 ha) of the natural landscape of peninsular Flor- ida in pre-Columbian times (fig. 1) (Auffenberg and Franz 1982). This xeric pineland occupies rolling to- pography of several ridge systems that run north-south, notably Trail Ridge, the Lake Wales Ridge, and the Brooksville Ridge; numerous lesser ridges and hills are identified by White (1970). These ridges consist of deep, well-drained soils of the Lakeland, Eustis, and Blanton asso- ciations (Beckenbach and Hammett 1962). Laessle (1942,1958a) describes the LLP/TO plant association as a fire climax system dominated by longleaf pine; slash pine (Pinus elli- otti) replaces longleaf pine in the community in south Florida. Turkey oak is a minor tree, but can achieve

co-dominance when fires are sup- pressed. The predominant under- story plant is wiregrass, Aristida stricta; however, a rich assemblage of perennial herbs vary in prominence in concert with seasonal changes. Monk (1968) recognizes two addi- tional phases of sandhill vegetation in north central Florida: (I) longleaf pine/sand post oak (Quercus nrarga- retta) and (2) longleaf pine/southern red oak (Quercus falcata). A fourth phase, longleaf pine/scrub hickory (Ca y a floridam), occurs in the south- ern portion of the Lake Wales Ridge (Abrahamson et al. 1984). Veno (1976), Givens et al. (1984), and Abra- hamson et al. (1984) provide quanti- tative data on LLP/TO community structure and dynamics. Myers (1985) suggests that longleaf pine/ turkey oak and sand pine scrub asso- ciations are successionally linked in some portions of their geographic ranges. Differences in physical / chemical features of soils of LLP/TO and SPS communities in the Ocala National Forest are not considered to be sufficient to explain the local dis-

Figure 1 .-Potential geographic distribution of longleaf pine/turkey oak sandhill and sand pine scrub xeric pinelands in Florida. Light shading indicates the sandhills and darker shading indicates the scrub. These distributions are based on Davis (1 980) and do not reflect minor sites of either commu- nity due to the scale of the itlustration.

tribu tion of the communities (Kalisz and Stone 1984).

Prior to settlement by European man, ground fires occurred in LLP/ TO sandhills at intervals of 1-5 years. These relatively "cool" fires favor regeneration of longleaf pine, flower- ing by grasses and herbs, and sup- press growth of woody plants (Myers 1985).

include those of Abrahamson et al. (1984) and Latham (1985).

Outstanding examples of SPS in- clude the "Big Scrub," part of the Ocala National Forest, scrubs of the Lake Wales Ridge, eg., the Archbold Biological Station, and stands along the Atlantic Coastal Ridge.

SMALL VERTEBRATE SPECIES ASSEMBLAGES

Sand Pine Scrub

Compared with the LLP/TO sandhill association, sand pine scrub (SPS) has less area (250,000 ha) and a far more limited distribution (fig. 1). Scrub is associated with old shore- lines, lake margins, and stream courses where extremely well washed, nutrient deficient sands were deposited during Pleistocene times (Kurz 1942; Laessle 1958a, b, 1967). The most widespread soils supporting SPS are the St. Lucie, Lakewood, and Pomello associations (Beckenbach and Hammett 1962).

Sand pine scrub is a two-layered community. Sand pine (Pinus clausa) normally occurs as a relatively even- aged overstory species. The under- story is comprised of 10-20 species of evergreen shrubs 1-5m in height. Four species of oaks comprise the bulk of the biomass, Quercus gemi- nata, Q. myrtifolia, Q. chapmanii, and Q. inapina. Lesser numbers of other species including Ceratiola ericoides, Lyonia ferruginea, and Osmanthus americanus add to local diversity. Sand pine scrub is a fire climax com- munity (Laessle 1958a, Abrahamson et al. 1984). In contrast with LLP/TO, SPS burns at intervals of 20-70 years; a combination of ground and crown fires destroys all the above-ground vegetation. Most of the woody plants, with the notable exception of the sand pine and Ceratiala, readily sprout from root crowns following fires. Laessle (1958a), Veno (1976), and Richardson (1 977) provide data on plant community structure of scrubs. Recent quantitative studies

Longleaf Pinepurkey Oak Sandhills

Amphibians and Reptiles

At least 47 species of herptiles, in- cluding 2 newts, 13 toads and frogs, 3 turtles, 10 lizards, 1 amphis- baenian, and 18 snakes, are reported to occur in LLP/TO habitats (table 1). Campbell and Christrnan (1982) list 5 categories of reptile and amphibian species that occur in LLP/TO and SPS: (1) characteristic (18 species); (2) associated with tortoise burrows (3 species); (3) frequent (8 species); (4) occasional (14 species); and (5) asso- ciated with aquatic habitats (21 spe- cies). Of the characteristic species, 7 are regarded as adapted to xeric con- ditions, 3 as sand swimmers, viz., Neoseps reynoldsi; Eumeces egregius, and Tantilla relicta, and the remainder (Sceloporus woodi, Mastico- phis flagellum, Stilosoma extenuaturn, Cnemidophorus sexlineatus) to other physical features of the habitats.

The gopher tortoise (Gopherus pol- yphemus) is a terrestrial turtle that digs deep burrows in the well- drained sandhill soils (Auffenberg and Franz 1982). Stout (1981) and Eisenberg (1983) recognized the go- pher tortoise was the keystone spe- cies in xeric pinelands. Some 80 spe- cies of animals may be classified as burrow commensals (Cox et al. 1987); however, the number of obligatory commensals is much smaller. Herp- tiles particularly associated with go- pher tortoise burrows include Rana

areolata, Pituophis melanoleucus, and D ymarchon corais.

The snake fauna of LLP/TO sandhills is species rich (L 18 spe- cies). This diversity includes large forms, e.g., D ymarchon corais couperi and Crotalus adamanteus, and small, specialized species like Stilosoma ex- tenuatum. This latter ophiophagous species feeds largely on Tantilla relicta; Tantilla, is in turn specialized on Tenebrionidae larvae (Mushinsky 1984).

Small Mammals

At least 19 species of small mammals with body masses less than 6.0 kg

may be anticipated in LLP/TO sand- hills (table 2). Two are fossorial, Scalopus aquatious and Geomys pinetia, 1 semi-fossorial, P. polionotus, and 2 occur in the surface litter, Blarina carolinensis and C yptotis parua.

Arboreal species include Sciurus carolinensis, S. niger, Glaucomys volans, P. gossypinus, and Ochrotomys nuttalli. Podomys floridanus nests in the burrows of the gopher tortoise and the pocket gopher (Layne 1969); it may enlarge other openings in the soil to establish burrows independ- ently of the gopher tortoise (R. E. Roberts, personal observation). Dasypus novemcinctus is the only ex- otic species of mammal that is clearly

established in the sandhill commu- ni ty.

Sand Pine Scrub

Amphibians and Reptiles

Campbell and Christman (1982) listed 64 species of reptiles and am- phibians that may be found in LLP/ TO sandhills and SPS. Pitfall trap- ping in six different even-aged stands of SPS on the Ocala National Forest by Christman et al. (unpub- lished manuscript and personal com- munication) revealed 27 species (table 1 ). Of 1,624 individuals

trapped, the common species were Bufo terrestris (n=332), Cnenridophorus sexlineatus (n=329), and Sceloporus woodi (n=216); five species were rep- resented by single captures. Christ- man et al. concluded that the herpe- tofaunal diversity declined with in- creasing age of SPS stands.

Copherus polyphemus is the key- stone species in SPS but is less com- mon there than in LLP/TO (Auffen- berg and Franz 1982). Many, if not most, of the burrow commensals are common in SPS (Cox et al. 1987). Podomys floridanus is an example.

Small Mammals

Fourteen species of small mammals commonly inhabit SPS (table 2). Pod-

omys floridanus is a predictable mem- ber of the assemblage throughout the range of scrubs in peninsular Florida (Layne 1978). Three subspecies of Peromyscus polionotus occur in scrubs of the interior and east coast portions of the peninsula. Common small mammals in central peninsular Flor- ida scrubs include Podomys floridanus, Perom yscus goss ypinus, Ochrotomys nuttalli, and Glaucomys volans (Swin- dell 1987). Podomys floridanus is the predominate small mammal in scrubs of southeast Florida (Richardson et al. 1986).

Limited data suggest Spilogale putorius is a major predator on small mammals in scrubs with lesser roles played by Mephitis mephitis and Mustela frenata (Stout and Roberts, personal observations).

ENDANGERED AND THREATENED SPECIES

Ten species of amphibians, reptiles, and small mammals associated with xeric pineland are currently listed as having some level of threatened, en- dangered, or sensitive status by ei- ther the state of Florida or the De- partment of Interior (table 3). The extensive overlap in species composi- tion between the two pineland com- munities results from the high num- ber of species common to both types. The Endangered Species Act charges federal agencies with the responsibil- ity to manage federally listed species on federally owned lands. At the state level, preservation of these listed species is of major concern when they occur on parcels of land scheduled for large-scale develop- ment. Preserve design and manage- ment practices for these species have largely evolved on an ad hoc basis without adequate time for an evalu- ation of the management or the long- term implications for the species.

MANAGEMENT OF XERIC PINELANDS ON PUBLIC LANDS

Of three national forests in Florida, only the Ocala National Forest is lo- cated in the peninsula. It totals 153,846 ha of which 85,020 ha are SPS and 18,219 ha LLP/TO. The National Forest Management Act (1976) and pursuant regulations (36 CFR 219) require that each forest be managed to maintain well-distributed and vi- able populations of wildlife species, including species that are endan- gered or threatened (Norse et al. 1986).

Silvicultural systems differ be- tween the two pineland communi- ties. On the Ocala National Forest sand pine scrub is routinely har- vested in patchy clearcuts that range from 16-24 ha in area. Scrub under- story vegetation is allowed to regen- erate naturally; however, sand pine is seeded following site preparation

by a single roller chopping. The har- vest rotation length is about 50 years. In contrast, LLP/TO is ostensibly managed on a 80-1 00 year rotation and shelterwood cutting favors natu- ral regeneration of the longleaf pine (Don Bethancourt, personal commu- nication). In practice, harvesting of longleaf pine may occur in 60 years.

Effectiveness of ecosystem man- agement in the SPS community will be judged by the response of desig- nated indicator species, such as go- pher tortoises and scrub jays Uphelo- coma coerulescens) (table 3). The go- pher tortoise is also a designated in- dicator species for the LEP/TO com- munity. The significance of the go- pher tortoise as a keystone species was emphasized in 1986 when har- vesting of the species on national for- ests in Florida was made illegal through an agreement between the

1 U.S. Forest Service and the Florida Game and Fresh Water Fish Csmmis- sion. Other species-specific manage- ment practices involving amphibians, reptiles, or small mammals have not

been deemed necessary to carry out on the Ocala National Forest (Don Bethancourt, personal communica- tion). In fact, the impact of timber harvesting on small vertebrates of LLP/TO and SPS communities is simply not known.

Public lands in Florida supporting xeric pinelands include, hut are not limited to, state forests and state parks. State forests with large acre- ages of LLP/TO, e.g., the Withla- coochee State Forest, are managed at the ecosystem level. Prescribed burn- ing is done every 3-8 years and fu- ture timber sales will follow a rota- tion length of 80-120 years; currently rotation lengths are about 60 years and are not regarded as favorably for endemic wildlife. Wildlife manage- ment areas overlap the state forest holdings and are managed for sus- tained yields of wildlife by the Flor- ida Game and Fresh h7ater Fish Commission based on a memoran- dum of understanding between agencies (Cathy Ryan, personal com- munication).

State parks are managed by the Division of Recreation and Parks of the Florida Department sf Natural Resources (FDNR). An ecosystem approach is taken in the restoration and management of xeric pinelands on state park lands (Jim Stevenson, personal communication). Prescribed burning has been used since 1969 to control hardwood invasion of LLP/ TO stands and to stimulate growth and flowering of grasses and herbs. Burning in spring and early summer appears to best duplicate the historic timing of lightning initiated fires in xeric pinelands. The impact of these management practices on the plant community has been documented (Davis 1984); the response of reptiles, amphibians, and small mammals is currently under study (Stout et al. unpublished). Generally, mature stands of SPS have not been burned until recently, due to the unpredict- able behavior of fire in the commu- nity; however, a prescription for burning this fuel type has been writ- ten and tested on private land and state parks (Doran et al. 1987). Early recovery stages of SPS appear to sup- port the greatest diversity of reptiles and amphibians. However, as can- opy closure occurs in SPS, ground cover diminishes and habitat quality for gopher tortoises declines (Cox et al. 1987). In contrast, similar numbers of Podomys have been observed in early (R. E. Roberts, unpublished data, J. Dickinson State Park); inter- mediate (Stout 1982); and old growth SPS (James N. Layne, unpublished data, Archbold Biological Station).

State parks, reserves, and pre- serves appear to be ideal lands to ex- plore species-specific management measures for herptiles and small mammals. For example, sand swim- ming herptiles (Smith 1982) require openings that are relatively root free in LLP/TO and SPS habitats. The natural occurrence of such openings may have been due to "hot" spots associated with the combustion of high fuel loads, e.g., fallen trees (Ron Myers, personal communication).

Concentration of natural fuels prior to prescription burns in SPS would offer a means to create microhabitat conditions favorable for the sand swimmers.

MANAGEMENT OF XERIC PINELAND ON PRIVATE LAND

Development of Regional Impact

Concern with management of am- phibians, reptiles, and small mam- mals on private lands in Florida de- rives from state and federal protec- tion of endangered species and the development guidelines promul- gated during the Development of Regional Impact (DRI) process. "The Florida Environmental Land and Water Management Act of 1972" (Chapter 380, Florida Statutes) de- fines developments of regional im- pact in Section 380.06(1), Florida Stat- utes, as "...any development which, because of its character, magnitude, or location, would have a substantial effect upon the health, safety, or wel- fare of citizens of more than one county (Anonymous 1976)." Large scale development projects in penin- sular Florida commonly involve hun- dreds to several thousand acres of relatively natural landscape. The DRI process requires bona fide studies of wildlife populations and their associ- ated habitats; emphasis is placed on listed species. Developers must pre- pare viable management strategies to accommodate wildlife resources de- pendent upon their lands (Cox et al. 1987; Richardson et al. 1986).

Management strategies of devel- opers with xeric pinelands generally follow one of two somewhat overlap- ping approaches to preserve habitat and/or species values: (1) conserva- tion set asides or (2) mitigation. Con- servation set asides are, in principle, the preferred solution. In practice some habitat is dedicated in perpetu- ity as a nature preserve; preserve de- sign currently is a somewhat ad hoc process and will be discussed more

completely in a subsequent section of this paper. Very high land values may dictate mitigation rather than on site preservation of habitat.

Mitigation may take many forms to compensate for development of xeric pinelands. Restoration of de- graded land (Humphrey et al. 19851, not necessarily xeric pinelands, is one method. Another tactic is to purchase comparable land or some other type of land of equivalent natural value elsewhere and dedicate it to preser- vation. A formal process for accom- plishing this option is presently un- der study by the Florida Game and Fresh Water Fish Commission.

Preservation of habitat is the basic purpose of conservation set asides and mitigations. The value of these efforts depends on the proximity to larger, undeveloped tracts of land, travel corridors, area of preserves, and future management options.

Another form of mitigation is the relocation of sensitive species from tracts of land to be developed to land dedicated to purposes that are con- sistent with the long-term survival of the relocated species. In Florida, the gopher tortoise has been the focus of numerous relocation efforts. Diemer (1984) discussed the advantages and disadvantages of relocation of go- pher tortoises as a species manage- ment strategy. Formal research on gopher tortoise relocation was re- cently reported (Proced. Gopher Tor- toise Relocation Symp., 27 June 1987, Gainesville, FL, in press). The Florida Game and Fresh Water Fish Commis- sion regulates relocations by a permit system based on a standardized relo- cation protocol.

Preserve Design

Preserve design is an evolving and controversial area of conservation biology (Diamond 1975,1978; Gilbert 1980; Higgs 1981; Margules 1982; Pickett and Thompson 1978; Pyle 1980; Soule and Simberloff 1986). Large preserves encompassing a mo-

saic of xeric pinelands, mesic forests, and seasonal and permanent wet- lands would perhaps offer the ideal landscape unit for long-term preser- vation of amphibians, reptiles, and small mammals in peninsular Flor- ida. Because preserves on private lands must be justified and dedicated through the DRI process, economics dictates preserve units of minimal size. Rarely do we have the opportu- nity to cluster or juxtapose these small units to take advantage of the so called "rescue effect" (Brown and Kodric-Brown 1977).

In practice, conservation set asides tend not only to be small in acreage but also only of one habitat type. The latter presents a dilemma for species whose requirements often include two or more contrasting habitats. For example, the gopher frog lives in tor- toise burrows in LLP/TO sandhills during late spring, summer and early fall and migrates to temporary wet season depressions to breed in win- ter and early spring (Moler and Franz 1987). Thus a mosaic of up- land-wetland habitats in close prox- imity are essential to maintain viable populations of this species. Other species such as the indigo snake have home range requirements that in- clude 122-202 ha of several upland- wetland habitat types (Moler 1985; Moler unpublished data). It is obvi- ous that large landscape units are necessary to preserve viable popula- tions of these animals.

We have prepared a detailed pre- serve design for a SPS community within the city of Boca Raton, Florida (Richardson et al. 1986; Stout et al. 1987; manuscript in preparation). The approach taken anticipated Soule and Simberloff (1986) and rec- ommended the area of the preserve be sufficient to support a minimum viable population (Franklin 1980) of gopher tortoises because of their status as the keystone species. Al- though biologically reasonable, this basis-for determining preserve size is often economically unrealistic from the view point of the private land-

owner. A consortium of public land- owners would, however, permit the purchase and long-term management of the preserve as recommended.

Cox et al. (1987) offer guidelines for the design of preserves on private lands to maintain gopher tortoise populations. They employed the computer simulation model POPDYN (Perez-Trejo and Samson manuscript) to determine population viability based on different initial sizes. Populations of 40-50 individu- als were found to be likely (290%) to persist 200 years. Based on existing literature on home range require- ments, Cox et al. (1987) recom- mended a minimum preserve of 10-

20 ha, depending on habitat quality, to support 40-50 tortoises.

Another approach to determining the area of a preserve employs "inci- dence functions" (Diamond 1978). Incidence functions are species spe- cific and derived from data sets which reveal the fraction of plots (discrete habitats) of different areas that actually support the species. It is a matter of judgement as to the probability of occurrence, e.g. 0.5 as opposed to 0.7, that would set a lower limit to area for an acceptable preserve. Data sets useful for evalu- ating this approach with respect to amphibians, reptiles, and small mammals in xeric pinelands are pres-

Cedar Grove 21.5 + + J. Dickinson 256.2 + +

ently lacking. Table 4 provides data we have gathered on area of discrete habitats and the presence or absence of gopher tortoises and Florida mice. It is apparent that tortoises are less area sensitive than Florida mice and that Florida mice are patchy in occur- rence in LLP/TO, perhaps only sec- ondarily related to area.

Incidence functions do not neces- sarily reveal the minimum area re- quired to support minimum viable populations (Franklin 1980). We be- lieve preserve area should be based on providing this requirement, par- ticularly when preserves are isolated relative to average dispersal dis- tances of keystone species. However, clusters of preserves within dispersal distances of keystone species may be of less area per preserve due to a high likelihood of reinvasion from nearby populations following local population extirpations (Noss and Harris 1986).

Management of Preserves in Xeric Pinelands

The future viability of preserves de- pends largely on their ownership af- ter development of the surrounding landscape. It is unlikely that home- owners associations will assume the cost of management if preserves re- main as a part of the overall develop- ment's "commons." Public owner- ship is an alternative and might rest with a city, county, or state. Local governments seem more appropriate; however, funds and expertise to manage may be lacking. One pre- serve in south Florida is designed to border a city park, thus allowing its maintenance and /or management costs to be assumed over time as part of the existing park system (Richardson, personal observation). Regardless of the ownership, a com- mi tment to long-term management must be achieved if a preserve is to retain natural values.

Management options for nature preserves range from a decision I ) to

do nothing and let nature take its course; 2) to manage for maintenance of a viable ecosystem, which implies the natural biota, including amphibi- ans, reptiles, and small mammals, will be present in proportion to their normal abundance; or 3) to focus management on the needs of one or more species. White and Bratton (1980) have exposed the folly of the first management option. The deci- sion to emphasize ecosystem or spe- cies management depends on the en- tity responsible for management, type of preserve, management objec- tives, area of the preserve, nature of the surrounding lands, relative over- all or regional rarity of particular species, and the resources available for management.

Management objectives of any preserve should focus on: I) mainte- nance of normal ecosystem proc- esses; 2) conservation of soil; 3) maintenance or restoration of normal hydrologic conditions; 4) prevention of establishment of exotic species.; 5) and prevention of human encroach- ment (e.g., dumping, ATVs, etc.) Be- yond these generalities, management of preserves is an idiosyncratic proc- ess that may concern endemic spe- cies, genetics of inbred populations, or restoration of periodic wild fires.

Xeric pinelands of peninsular Flor- ida depend on periodic fires to main- tain their structure and function (Laessle 1958a; Abrahamson 1984). Thus a burning program is essential in the management of LLP/TO or SPS preserves. Spring or early sum- mer prescribed burns are routinely used to maintain LLP/TO communi- ties on state parks. Doran et al. (1 987) have documented prescribed burns of SPS preserves in an urban setting based on rather esoteric fire models developed by the U.S. Forest Service. ~ o ~ h e r tortoises respond favorably to the bums (Stout et al. 1988). A mo- saic of recovery stages in SPS may favor beta diversity of herptiles and small mammals. Mushinsky (1985) has carefully documented the re- sponse of the herpetofauna to a vari-

ety of burning schedules in LLP/TO. Diversity and abundance of amphibi- ans and reptiles was increased on experimental plots relative to un- burned controls. Re-establishment of the pine overstory may be necessary to produce needle cast for carrying fire (Landers and Speake 1980).

Management of conservation set a s id s and/or easements may focus on particular species or combinations of species. The smaller the preserve the more likely that a reduced suite of species will be present (Richardson et al. 1986). Given that a fixed area is available for manage- ment, major efforts to enhance or maintain habitat should target those species that can maintain viable gopula tions within the preserve (Shaffer 1986). A species whose mini- mum area requirements for a mini- mum viable population exceeds the preserve area should not be of major concern (Shaffer and Samson 1985); nonetheless, such species can benefit from the preserves if travel corridors exist (Harris 1984).

DISCUSSION

Xeric pinelands of peninsular Florida support a species-rich assemblage of reptiles, amphibians, and small mammals. Growth and development continues to diminish LLP/TO and SPS habitats to the detriment of the associated biota. Land in public own- ership, e.g. state parks and forests, national forests, and private hold- ings, e.g., the Archbold Biological Station, and institutional lands such as the Ordway and Swisher Pre- serves, jointly owned and managed by the University of Florida and The Nature Conservancy, will be increas- ingly valuable as other xeric pine- lands are converted to land uses not favorable to the biota. Thus, manage- ment of these xeric pinelands will become more important in the fu- ture. At present management is largely limited to prescribed bums to maintain what were historically fire

climax communities. Thus, fire man- agement is tantamount to small ver- tebrate management.

In the future as air quality stan- dards are modified, prescribed burn- ing, particularly in or near urbanized areas, will be restricted or eliminated as a management option. Alternative means of habitat manipulation need to be developed, particularly for SPS.

Basic information on the life his- tory of many amphibians, reptiles, and small mammals of xeric pine- lands is lacking. The Nongarne Wild- life Program of the Florida Game and Fresh Water Fish Commission has initiated and f u ~ ~ d e d rather large scale studies of SPS and LLP/TO communities. These studies are at the community level and largely obser- vational. Management needs of indi- vidual species may be derived only secondarily from this research. Stud- ies that focus on particular species will ultimately lead to more refined habitat management guidelines. The report by Cox et al. (1987) will likely serve as a model for the preparation of habitat protection guidelines; man- agement follows protection (White and Bratton 1980).

Management alternatives at the ecosystem and species level are needed now for xeric pinelands on private lands undergoing develop- ment Regulation of development in these habitats as currently practiced will result in a patchwork of small, isolated nature preserves. Preserva- tion of natural habitat in a developed landscape is, of course, desirable. However, several problems remain: (1) who will own the preserves, (2) how will a management plan be pre- pared, and (3) who will be respon- sible for management? Even another decade of rapid growth in peninsular Florida may result in a feG hundred nature preserves, which will not nec- essarily be restricted to xeric pine- land habitat. Ignoring the question of ownership, no public land manage- ment agency is currently capable of assuming the charge of managing these preserves. Lack of manage-

ment, e.g., failure to conduct pre- scribed burning, will allow succes- sional changes to occur to the detri- ment of many small vertebrates nar- rowly adapted to xeric pinelands. Loss of habitat and species values originally used by jurisdictional agencies to secure preserve set asides provides a potential basis for private land owners to request development rights on the land. This action would defeat the entire purpose of having conservation set asides.

An alternative to on site habitat protection is offered by Cox et al. (1987) in regard to preserving habitat for the gopher tortoise. The alterna- tive, a Wildlife Resource Mitigation Fund (WRMF), allows a developer to contribute money to the fund to miti- gate losses of valuable wildlife habi- tat on lands being developed. The collective monies of several develop- ment projects would allow an inde- pendent group such as the Trust For Public Lands to assist in the purchase of commensurate lands to expand an existing public park, preserve or for- est. Management is more likely to be applied to these lands and ultimately the resources are better served by the public agencies.

ACKNOWLEDGMENTS

We thank the authors of the papers cited herein for their efforts and dedication to science. Biologists who contributed to our knowledge of xeric pineland include but are not limited to the following individuals: Dan Austin, Don Bethancourt, Russ Burke, Steve Christman, David Cook, David Corey, Jim Cox, Joan Diemer, Dick Franz, Larry Harris, Randy Kautz, Jim Layne, Wayne Marion, Paul Moler, Ron Myers, Reed Noss, Cathy Ryan, Jim Stevenson, and Don Wood. Support for research on the ecology of sandhill communities was provided by the Nongame Wildlife Program, RFP86-003, of the Florida Game and Fresh Water Fish Commis- sion. Development groups that

funded work by the authors on xeric pinelands include Hardy-Lieb Devel- opment Corporation, The Adler Group, and Deutsch-Ireland Proper- ties. The Division of Recreation and Parks (FDNR), Department of Biol- ogy, University of South Florida, and the Department of Biological Sci- ences, University of Central Florida assisted in our studies in a variety of ways. We thank Beverly Bamekow, Rita Greenwell, Barbara Erwin, and Nancy Small for typing the manu- script. Lastly, we thank Paul E. Moler, James N. Layne and Robert C. Szaro for providing excellent sugges- tions to improve the paper.

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